PHOTOSYNTHETIC PHYSIOLOGY OF PROROCENTRUM MARIAE-LEBOURIAE (DINOPHYCEAE) DURING ITS SUBPYCNOCLINE TRANSPORT IN CHESAPEAKE BAY1

This paper presents results of field studies on the estuarine dinoflagellate Prorocentrum mariae-lebouriae (Parke & Ballantine) Faust in Chesapeake Bay. We tested the hypothesis that the photosynthetic physiology of Prorocentrum shows adaptive responses to low-light during a lengthy subpycnocline transport in estuarine circulation. Prorocentrum underwent a seasonal, northward trnasport between February and June, 1984 and 1985. Low cell densities occurred in the seaward part of the estuary during winter and early-spring, subpycnocline populations progressed up-estuary in the ensuring 2–3 months, and dense surface populations developed in the mesohaline portion of the estuary thereafter. We sampled Prorocentrum from surface and subpycnocline waters and measured photosynthesis-light (P-I) relations with in situ incubations. The photophysiology of Prorocentrum collected below the pycnoline differed from that of cells in the surface mixed layer in that photosynthetic efficiency, α-cell^?1, was higher, photosynthetic capacity, P_max-cell^?1 was ·4 times greater for subpycnocline (≦ 10m) samples than for those from the surface mixed layer (≧ 6m). Comparison of in situ photosynthetic properties to those generated in laboratory studies showed that values of α·cell^?1 for both surface and subpycnocline samples were in the range found for cultures in low-light. Concentrations of Chls a, c and peridinin·cell^?1 and molar pigment ratios peridinin: Chl a and Chl a: Chl c were not significantly different for the surface and subpycnocline samples, nor were C · cell^?1 or C : Chl a. Chloroplast and starch volume fractions and the number of thylakoids were the same for samples collected at different depths, and there was no evidence of cytoplasmic vacuolization in any field samples. These morphometric data for cells from natural populations of Prorocentrum most closely resembled data for laboratory cultures grown at or near 2.6E·m-^?2·4d^?1. A lower growth irradiance of 0.3E·m^?2·d^?1 produced indications of stress in cultures, including starch depletion and vacuolization, that were never observed in natural populations. Based on the combination of these findings, we conclude that Prorocentrum is adapted to low-light both in the surface mixed layer and beneath the pycnocline, although certain photophysiological characteristics distinguish these two groups of samples.     

THE TIME-COURSE OF PHOTOADAPTATION TO LOW-LIGHT IN PROROCENTRUM MARIAE-LEBOURIAE (DINOPHYCEAE)1

The estuarine dinoflagellate, Prorocentrum mariaelebouriae (Parke & Ballantine 1957) Faust 1974 undergoes increases in pigmentation and photosynthetic efficiency within several days of downward light shifts. These changes can be described by first-order kinetics, as has been reported previously for Chlorophyll (Chl) a in several phytoplankton species. The studies described in this paper were conducted with isolates of populations of Prorocentrum from the Chesapeake Bay. We determined rates of adaptation to low-light for cultures grown at a range of photon flux densities (I₀= 2.65–26.2 E.m^?2, d^?1, shifted to 6.3–7.0% I₀) at three temperatures (10°, 15°, and 20° C), bracketing the conditions this species experiences in situ. In this paper, I report the time-course of changes in α, P_max Chl a, peridinin, and I_k and first-order rate constants, K₁ for changes in α, Chl a and peridinin. cell^?1. K₁ for changes in α cell^?1 averaged 1.58 × 10^?2 h^?1 for conditions encompassing five light treatments and three temperatures; the corresponding mean for Chl a was 1.59 × 10^?2 h^?1. Increases in peridinin measured for five light treatments at 15° C showed a mean K₁ of 1.22 × 10^?2 h^?1, Average percent changes in per cell α, Chl a, and peridinin ranged from 0.4–4.0% h^?1 (10–90% d^?1) following exposure to low-light. Photoadaptive changes are important to Prorocentrum because in nature it occupies turbid waters (K_t≥ 0.5 m^?1) where the mixing depth often exceeds the depth of the photic layer. Cells are entrained beneath a seasonally-stable density discontinuity and are exposed to very low-light (< I E.m^?2.d^?1) for days to weeks during subpycnocline transport. The ability of this species to undergo changes in pigmentation and photosynthetic physiology confers increased efficiency of light harvesting and contributes to this species’survival in the estuary where it is an important component of the dinoflagellate flora.     

Deep-water aquatic plant communities in an oligotrophic lake: physiological responses to variable light

1. Oligotrophic Lake Waikaremoana, New Zealand, is used for hydroelectric power generation and the lake levels are manipulated within an operating range of 3 m. There was concern that rapidly changing water levels adversely affected the littoral zone by decreasing light availability in two ways: local turbidity caused by shoreline erosion at low water levels; and decreased light penetration to the deep littoral zone caused by high water levels in summer. 2. The littoral zone was dominated by native aquatic plants with vascular species to 6 m and a characean meadow below this to 16 m. The biomass and heights of the communities in the depth zone 0–6 m were reduced at a site exposed to wave action relative to those at a sheltered site. However, the community structure below 6 m was similar at exposed and sheltered sites. The lower boundary of the littoral zone was sharply delimited at 16 m and this bottom boundary remained constant throughout the year despite large seasonal changes in solar radiation and the 3 m variation in lake level. 3. There was evidence that the deep-water community consisting of Chara corallina had adapted physiologically to low-light conditions. Net light saturated photosynthesis (CO₂ exchange) per unit chlorophyll a (Chl a) was reduced to 1.7 μg C (μg Chl a)^?1 h^?1 at the lower boundary, half of that recorded at 5 m. The concentration of Chi a per gram of biomass (dry weight), was considerably greater at the lower boundary than higher in the profile [c. 7 mg Chl a (g dry wt)^?1 at 16 m vs. 4 mg Chl a (g dry wt)^?1 at 5 m]. Chl b also increased with depth and there was no change in the ratio of Chl a and Chl b with increasing depth. The saturation light intensity (I_k) of the community at the lower boundary was only 78 μmol photons m^?2 s^?1. Photosynthetic parameters (I_k and α) as well as the Chl a content remained relatively constant throughout the seasonal and short-term changes in radiation. 4. The photosynthetic characteristics of the littoral community were therefore not greatly affected by the lake level change caused by the present hydroelectric operations. However, the sharpness of the lower boundary and its extreme shade characteristics imply that the deep-water community would be sensitive to any further changes in underwater light availability.     

Electron Microscopic Observations on the Symbiosis of Paramecium bursaria and its Intracellular Algae*

SYNOPSIS. Observations were made on the fine structure of Paramecium bursaria and its intracellular Chlorella symbionts. Emphasis was placed on the structure of the algae and structural aspects of the relationship between the organisms. The algae are surrounded by a prominent cell wall and contain a cup-shaped chloroplast which lies just beneath the plasma membrane. Within the cavity formed by the chloroplast are a large nucleus, a mitochondrion, one or more dictyosomes, and numerous ribosomes. The chloroplast itself is made up of a series of lamellar stacks each containing 2–6 or more thylakoids with a granular stroma and starch grains intercalated between the stacks. The thylakoid stacks of mature algae are frequently more compact than those of recently divided algae. A large pyrenoid is located within the base of the chloroplast. It is made up of a granular or fibrillar matrix surrounded by a shell of starch. The matrix is bisected by a stack of 2 thylakoids. Prior to the division of the chloroplast the pyrenoid regresses; pyrenoids subsequently form in the daughter chloroplasts thru condensation of the matrix material and the reappearance of a starch shell. This shell appears to be formed by the hollowing-out of starch grains already present in the chloroplast stroma. Accordingly, in this case, starch moves from the stroma to the pyrenoid. The algae are located thruout the peripheral cytoplasm of the Paramecium. Each alga is located in an individual vacuole except immediately following division of the algae when the daughter cells are temporarily located in the vacuole which harbored the parental cell. Shortly thereafter the vacuole membrane invaginates, thereby isolating the daughter algae into individual vacuoles. Degenerating symbiotic algae are seen; because these are frequently found in vacuoles with bacteria, they are presumed to be undergoing digestion. Due to the conditions of culture these algae could have been either of intracellular or extracellular origin.     

CHANGES IN THE ULTRASTRUCTURE OF SYMBIOTIC ZOOXANTHELLAE (SYMBIODINIUM SP., DINOPHYCEAE) IN FED AND STARVED SEA ANEMONES MAINTAINED UNDER HIGH AND LOW LIGHT1

The ultrastructure of symbiotic dinoflagellates (Symbiodinium sp., zooxanthellae) in the sea anemone Aiptasia pallida Verrill was examined in well-fed or starved (up to 120 days) anemones maintained under two light levels (5 and 50 μmol · m^?2· s^?1). Cell size of zooxanthellae was not affected by feeding history; however, both light and feeding history affected the relative cell volume of chloroplasts, lipids, and vacuoles. Stereological analysis of transmission electron micrographs showed that algae in low-light starved anemones had 10 times as much lipid (17.4% of cell volume) as those in well-fed anemones under the same light conditions (1.8%). The lipid content of algae from anemones in high light increased from 15.4% in well-fed anemones to 30.1% in starved anemones. The starch content of zooxanthellae in low-light anemones was law (4.1%) and not affected by feeding history, while the starch content of zooxanthellae in high-light anemones was greater (10.7%), with some differences among groups. Algal photoacclimation to low light included an increase in chloroplast relative volume from 17% (in well-fed high-light anemones) to 33% in well-fed low-light anemones. Starvation of the host resulted in a significant decrease in chloroplast volume in zooxanthellae in anemones at both light levels. Morphometry provides quantitative confirmation of biochemical and physiological data on zooxanthellae, because the changes in zooxanthellae with starvation of the host are consistent with other indicators of nutrient limitation of zooxanthellae of A. pallida held without food for long periods of time.     

Dunaliella salina (Chlorophyta) with small chlorophyll antenna sizes exhibit higher photosynthetic productivities and photon use efficiencies than normally pigmented cells

The photon use efficiencies and maximal rates of photosynthesis in Dunaliella salina (Chlorophyta) cultures acclimated to different light intensities were investigated. Batch cultures were grown to the mid-exponential phase under continuous low-light (LL: 100 μmol photon m^-2 s^-1) or high-light (HL: 2000 μmol photon m^-2 s^-1) conditions. Under LL, cells were normally pigmented (deep green) containing ∼500 chlorophyll (Chl) molecules per photosystem II (PSII) unit and ∼250 Chl molecules per photosystem I (PSI). HL-grown cells were yellow-green, contained only 60 Chl per PSII and 100 Chl per PSI and showed signs of chronic photoinhibition, i.e., accumulation of photodamaged PSII reaction centers in the chloroplast thylakoids. In LL-grown cells, photosynthesis saturated at ∼200 μmol photon m^-2 s^-1 with a rate (P_max) of ∼100 mmol O₂ (mol Chl)^-1 s^-1. In HL-grown cells, photosynthesis saturated at much higher light intensities, i.e. ∼2500 μmol photon m^-2 s^-1, and exhibited a three-fold higher P_max (∼300 mmol O₂ (mol Chl)^-1 s^-1) than the normally pigmented LL-grown cells. Recovery of the HL-grown cells from photoinhibition, occurring prior to a light-harvesting Chl antenna size increase, enhanced P_max to ∼675 mmol O₂ (mol Chl)^-1 s^-1. Extrapolation of these results to outdoor mass culture conditions suggested that algal strains with small Chl antenna size could exhibit 2–3 times higher productivities than currently achieved with normally pigmented cells. This revised version was published online in August 2006 with corrections to the Cover Date.     

Enhancement of susceptivity to photoinhibition and photooxidation in rice chlorophyll <Emphasis Type="Italic">b</Emphasis>-less mutants

Two rice chlorophyll (Chl) b-less mutants (VG28-1, VG30-5) and the respective wild type (WT) plant (cv. Zhonghua No. 11) were analyzed for the changes in Chl fluorescence parameters, xanthophyll cycle pool, and its de-epoxidation state under exposure to strong irradiance, SI (1 700 μmol m⁻² s⁻¹). We also examined alterations in the chloroplast ultrastructure of the mutants induced by methyl viologen (MV) photooxidation. During HI (0–3.5 h), the photoinactivation of photosystem 2 (PS2) appeared earlier and more severely in Chl b-less mutants than in the WT. The decreases in maximal photochemical efficiency of PS2 in the dark (F_v/F_m), quantum efficiency of PS2 electron transport (Φ_PS2), photochemical quenching (q_P), as well as rate of photochemistry (P_rate), and the increases in de-epoxidation state (DES) and rate of thermal dissipation of excitation energy (D_rate) were significantly greater in Chl b-mutants compared with the WT plant. A relatively larger xanthophyll pool and 78–83 % conversion of violaxanthin into antheraxanthin and zeaxanthin in the mutants after 3.5 h of HI was accompanied with a high ratio of inactive/total PS2 (0.55–0.73) and high 1–q_P (0.57–0.68) which showed that the activities of the xanthophyll cycle were probably insufficient to protect the photosynthetic apparatus against photoinhibition. No apparent difference of chloroplast ultrastructure was found between Chl b-less mutants and WT plants grown under low, LI (180 μmol m⁻² s⁻¹) and high, HI (700 μmol m⁻² s⁻¹) irradiance. However, swollen chloroplasts and slight dilation of thylakoids occurred in both mutants and the WT grown under LI followed by MV treatment. These typical symptoms of photooxidative damage were aggravated as plants were exposed to HI. Distorted and loose scattered thylakoids were observed in particular in the Chl b-less mutants. A greater extent of photoinhibition and photooxidation in these mutants indicated that the susceptibility to HI and oxidative stresses was enhanced in the photosynthetic apparatus without Chl b most likely as a consequence of a smaller antenna size.     

Photoacclimation characteristics of Sargassum thunbergii germlings under different light intensities

The photosynthesis and growth responses of Sargassum thunbergii germlings to different light intensities (10, 60, and 300 μmol photons m^?2 s^?1) were investigated. Maximum photochemical efficiency (F _v/F _m), rapid light curves (RLCs), and photochemical and non-photochemical quenching (qP and NPQ) were estimated by a pulse amplitude-modulated fluorometer. The photosynthesis of S. thunbergii germlings exhibited different properties to optimize light capture and utilization. The excitation pressure (1???qP) was rapidly increased to approximately 0.27 showing that germlings responded to high light by chronic photoinhibition with an accumulation of closed reaction centers, which ultimately resulted in a slow growth. This was accompanied by a reduced F _v/F _m with time and a development of high capacity for NPQ. Although F _v/F _m in moderate-light germlings did not fully recover overnight, germlings demonstrated a less severe chronic photoinhibition considering the reduced degree of excitation pressure accumulation of approximately 0.15. The relative stability of photosynthetic capacity (rETR_max, E _k, and α) could endow germlings with the highest relative growth rate (RGR) of approximately 9.3 % day^?1 in moderate light. By contrast, low-light germlings demonstrated high F _v/F _m and F _o, corresponding high α collectively suggested greater efficiency of light absorption and energy transformation. Sustained increases in electron transport capacity (rETR_max and E _k) occurred in low-light germlings, which resulted in a stable RGR of over 8.2 % day^?1. Consequently, S. thunbergii germlings are considered to prefer low light regimes and have a relative capacity of moderate and high light tolerance. However, the light acclimation to oversaturating conditions is at the cost of slow growth to maintain survival.     

Chloroplast position and photosynthetic characteristics in two monostromatic species,Monostroma angicava and Protomonostroma undulatum (Ulvophyceae), having a shared ecological niche

Monostroma angicava and Protomonostroma undulatum are monostromatic green benthic algae (Ulvophyceae), which grow together in the same intertidal habitat of Muroran, Hokkaido, Japan, during the spring season. Commonly, both species have a single chloroplast with one pyrenoid per cell. The parietal chloroplast is located on the periphery of the thallus in both species, although the location of the chloroplast differs in the two. In M. angicava , the chloroplast was observed to be arranged on one‐side of the thallus surface, whereas, in P. undulatum , it was dispersed and randomly located on either side of the thallus or on the lateral face. The density of chlorophylls (Chls) assessed from the absorption spectra of the thallus and its solvent extract was higher in M. angicava , which appeared dark‐green in color, than in the light‐green colored P. undulatum . The maximum photosynthetic rate per thallus area (μmol O₂ m^?2 s^?1) was higher in M. angicava , whereas, per total chlorophyll content (μmol O₂ g Chl a + Chl b ^?1 s^?1) was higher in P. undulatum . Both species showed similar efficiency of photosynthesis at light‐limiting conditions. The efficiency of light absorption by photosystem II (PSII ) in P. undulatum was higher than M. angicava , whereas the photoprotective response was higher in M. angicava . This indicates that more energy is utilized in M. angicava to protect its PSII due to the chloroplast position, which has more direct exposure to light and, therefore, lowers the efficiency of light absorption by PSII . The higher density of chlorophylls in M. angicava could explain higher photosynthesis per thallus area, whereas, higher efficiency of light absorption by PSII in P. undulatum could explain higher photosynthesis per total chlorophyll content. The differences in light absorption efficiency and quantum efficiency of PSII might be an important ecological strategy in these two species for their coexistence in the intertidal area.     

Disintegration of chloroplasts during zygote formation inSpirogyra verruculosa

The chloroplast disintegration during zygote maturation inSpirogyra verruculosa was investigated by electron microscopy. In the seven-day-old zygote about half of the chloroplasts commenced to disintegrate and to turn yellow, losing starch grains, and, then, were torn into fragments of various sizes, which had mostly vesiculated thylakoids and plastoglobules increasing in both size and number. At about two weeks after conjugation, in the cytoplasm, electron-dense structures, linear in section, appeared and vacuoles of various sizes developed. Each of the dense linear structures lying around a fragment seemed to form a cavity of crescent shape in section, and these cavities fused mutually into a large one, leading to the separation of the fragment from the bulk of cytoplasm. The vacuoles seemed to be, involved in the sequestration of the fragments by their fusion with the cavities and by the invagination, of tonoplast. The fragments entrapped by the vacuoles were rapidly broken down into the aggregation of residual membrane pieces, plastoglobules, and undigested starch grains. The maintained chloroplasts changed little in structure compared with the chloroplast of the vegetative cell, and were transmitted to the germling. It is suggested that the eliminated chloroplasts are derived exclusively from the male gamete.     

PRODUCTIVITY OF THE FILAMENTOUS ALGA PITHOPHORA OEDOGONIA (CHLOROPHYTA) IN SURREY LAKE,INDIANA1

Biomass, akinete numbers, net photosynthesis, and respiration of Pithophora oedogonia were monitored over two growing seasons in shallow Surrey Lake, Indiana. Low rates of photosynthesis occurred from late fall to early spring and increased to maximum levels in late spring to summer (29–39 mgO₂·g^?1 dry wt·h^?1). Areal biomass increased following the rise in photosynthesis and peaked in autumn (163–206g dry wt·m^?2). Photosynthetic rates were directly correlated with temperature, nitrogen, and phosphorus over the entire annual cycle and during the growing season. Differences in photosynthetic activity and biomass between the two growing seasons (1980 and 1981) were apparently related to higher, early spring temperatures and higher levels of NO₃-N and PO₄-P in 1981. Laboratory investigations of temperature and light effects on Pithophora photosynthesis and respiration indicated that these processes were severely inhibited below 15°C. The highest P_max value occurred at 35°C (0.602 μmol O₂·mg^?1 chl a·min^?1). Rates of dark respiration did not increase above 25°C thus contributing to a favorable balance of photosynthetic production to respiratory utilization at high temperatures. Light was most efficiently utilized at 15°C as indicated by minimum values of I_k(47 μE·m^?2·s^?1) and I_c (6 μE·m^?2·s^?1). Comparison of P. oedogonia and Cladophora glomerata indicated that the former was more tolerant of temperatures above 30°C. Pithophora's tolerance of high temperature and efficient use of low light intensity appear to be adaptive to conditions found within the dense, floating algal mats and the shallow littoral areas inhabited by this filamentous alga.     

Photoautotrophic potato cells: Transition from heterotrophic to autotrophic growth

Cells of potato (Solanum tuberosum L.) were obtained which were capable of photoautotrophic growth in liquid suspension culture under a photon flux density of 90–110 μmol m^?2 s^?1 PAR and in an atmosphere enriched with 2% CO₂. These photoautotrophic cells contained between 100 to 200 μg Chl (g fresh weight)^?1 and fixed CO₂ at a maximum rate of 16 μmol CO₂ (g fresh weight)^?1h^?1. In order to obtain cells capable of photoautotrophic growth it was necessary to adapt highly chlorophyllous heterotrophic cells (>50 μg Chl (g fresh weight)^?1) for growth in medium with 2.5 g sucrose 1^?1 (photomixotrophic cells). The photomixotropic cells had a Chl content of ca 100 μg Chl (g fresh weight)^?1 and were capable of photosynthetic activity which allowed them to survive after sugars had been depleted from the medium. It was from the photomixotrophic cells that cells capable of photoautotrophic growth were obtained. Heterotrophic cells initially established in liquid medium with 25 g sucrose I^?1 from chlorophyllous callus contained about 50 to 150 μg Chl (g fresh weight)^?1. However, after 5 to 10 passages the Chl content decreased to a maximum of 15 μg Chl (g fresh weight)^?1. These cells could not be adapted to photomixotrophic or photoautotrophic growth. These cells also were not able to regain Chl or initiate high rates of CO₂ fixation during the stationary phase of growth as did photomixotrophic cells or chlorophyllous heterotrophic cells. The loss of Chl exhibited by the cells during adaption to heterotrophic growth could be attributed at least in part to unbalanced growth (when cell division and growth exceeds Chl accumulation). Sucrose appeared to have an inhibitory effect directly on photosynthesis independent of Chl accumulation.     

dr and spr/sr mutations of Chlamydomonas reinhardtii affecting D1 protein function and synthesis define two independent steps leading to chronic photoinhibition and confer differential fitness

The effects of introduced chloroplast gene mutations affecting D1 synthesis, turnover and function on photosynthesis, growth and competitive ability were examined in autotrophic cultures of Chlamydomonas reinhardtii (Chlorophyta) adapted to low or high irradiance. Few discernible effects were evident when the mutants were grown in low light (LL, 70 μmol m^?2 s^?1). The herbicide-resistant psbA mutation Ser²⁶⁴→ Ala (dr) slowed electron transfer and accelerated D1 degradation in cells grown under high light (HL, 600 μmol m^?2 s^?1). The maximum rate of light-and CO₂-saturated photosynthesis, cell growth rate and competitive ability in the dr mutant were reduced compared to wild type under HL. However, the wild-type rate of D1 synthesis in dr was adequate to compensate for accelerated D1 degradation. 16S rRNA mutations conferring resistance to streptomycin and spectinomycin (spr/sr) that altered chloroplast ribosome structure and assembly were used to inhibit chloroplast protein synthesis. In spr/sr cells grown under HL, D1 synthesis was reduced by 40–60% compared to wild type and D1 degradation was accelerated, leading to a 4-fold reduction in D1 pool size. The reduced D1 levels were accompanied by an elevation of F_o and a decline in F_v/F_m, quantum yield and maximum rate of CO₂-saturated photosynthesis. Chemostat experiments showed that the growth rate and competitive ability of spr/sr were reduced against both wild type and dr.     

Photosynthetic apparatus organization and function in the wild type and a chlorophyll b-less mutant of Chlamydomonas reinhardtii. Dependence on carbon source

 The assembly, organization and function of the photosynthetic apparatus was investigated in the wild type and a chlorophyll (Chl) b-less mutant of the unicellular green alga Chlamydomonas reinhardtii, generated via DNA insertional mutagenesis. Comparative analyses were undertaken with cells grown photoheterotrophically (acetate), photomixotrophically (acetate and HCO⁻ ₃) or photoautotrophically (HCO⁻ ₃). It is shown that lack of Chl b diminished the photosystem-II (PSII) functional Chl antenna size from 320 Chl (a and b) to about 95 Chl a molecules. However, the functional Chl antenna size of PSI remained fairly constant at about 290 Chl molecules, independent of the presence of Chl b. Western blot and kinetic analyses suggested the presence of inner subunits of the Chl a-b light-harvesting complex of PSII (LHCII) and the entire complement of the Chl a-b light-harvesting complex of PSI (LHCI) in the mutant. It is concluded that Chl a can replace Chl b in the inner subunits of the LHCII and in the entire complement of the LHCI. Growth of cells on acetate as the sole carbon source imposes limitations in the photon-use efficiency and capacity of photosynthesis. These are manifested as a lower quantum yield and lower light-saturated rate of photosynthesis, and as lower variable to maximal (F_v/F_max) chlorophyll fluorescence yield ratios. This adverse effect probably originates because acetate shifts the oxidation-reduction state of the plastoquinone pool, and also because it causes a decrease in the amount and/or activity of Rubisco in the chloroplast. Such limitations are fully alleviated upon inclusion of an inorganic carbon source (e.g. bicarbonate) in the cell growth medium. Further, the work provides evidence to show that transformation of green algae can be used as a tool by which to generate mutants exhibiting a permanently truncated Chl antenna size and a higher (per Chl) photosynthetic productivity of the cells. Received: 10 November 1999 / Accepted: 22 December 1999     

NITROGEN LIMITATION IN ISOCHRYSIS GALBANA (HAPTOPHYCEAE). I. PHOTOSYNTHETIC ENERGY CONVERSION AND GROWTH EFFICIENCIES1

The effect of steady-state nitrogen limitation on photo-synthetic characteristics and growth efficiency was examined in the marine haptophyte Isochrysis galbana Green. Nitrate limited chemostats were maintained at nine dilution rates, ranging from 0.18-0.96 d^?1, under continuous irradiance levels of 175 μmole quanta·m^?2·s^?1, an irradiance level which saturated photosynthesis at all growth rates. Nitrogen limitation led to an overall reduction in pigmentation and a decrease in the cellular concentration of reaction centers; however, the optical absorption cross section, normalized to Chl a, increased. Moreover, Chl c/a ratios were higher in nitrogen-limited cells: the change in Chl c/a ratios were correlated with an increase in the functional size of Photosystem II. Both light saturated photosynthetic rates normalized per cell and specific respiratory losses were positively linearly correlated with growth rate. Light saturated photosynthetic rates normalized to Chl a remained relatively insensitive to the rate of nitrogen supply. The minimum quantum requirement for gross photosynthetic oxygen evolution increased from 12.4 to 17.0 quanta/O₂. At the growth irradiance, the quantum requirement increased 88%, from 19.9 to 37.5 quauta/O₂ Photosynthesis/respiration ratios remained relatively constant at dilution rates greater than 35% of the maximum relative growth rate. Consequently, net growth efficiency, defined as the ratio of the specific growth rate, μ, to specific gross photosynthesis, P, also remained relatively constant over this range of growth rates averaging 85 ± 3%.     

Light acclimation in leaves of the juvenile and adult life phases of ivy (Hedera helix)

Hoflacher, H. and Bauer, H. 1982. Light acclimation in leaves of the juvenile and adult life phases of ivy (Hedera helix). – Physiol. Plant. 56: 177–182. Light acclimation was investigated during the juvenile and adult life phases of the whole-plant-development in Hedera helix L. For this purpose, cuttings of the juvenile and adult parts of one single parent plant were grown under low-light (PAR 30–50 μmol photons m^?2 s^?1) and high-light (PAR 300–500 μmol m^?2 s^?1) conditions: CO₂ exchange, chloroplast functions, and specific anatomy of fully developed leaves differentiated under these conditions were determined. In juvenile plants the leaves formed under low and high light had light-saturated rates of net photosynthesis of 6.5 and 11.1 mg CO₂ (dm leaf area)^?2 h^?1, respectively. In adult plants the rates were 9.4 and 22.2 mg dm^?2 h^?1, indicating a more pronounced capacity for acclimation to strong light in the adult life phase. Higher photosynthetic capacities were accompanied by higher conductances for the CO₂ transfer through the stomata, leading to almost the same CO₂ concentration in the intercellular spaces. Thus, stomatal conductances were not primarily responsible for the different photo-synthetic capacities. The higher rates in adult and high-light grown leaves were mainly the result of formation of thicker leaves with more chloroplasts per unit leaf area. Expressed per chloroplast, the photosynthetic capacity, the Hill reaction, and the activity of ribulose bisphosphate carboxylase were almost identical in plants grown in low-light and high-light. Measurements of photosynthetic capacity and thickness of leaves of Hedera sampled from field habitats with contrasting light regimes confirm the results of growth chamber studies. It is, therefore, concluded that both life phases of Hedera are capable of acclimating to strong light, but that during the juvenile phase this capacity is not fully developed.     

PHOTOSYNTHESIS AND RESPIRATION OF THE CONCHOCELIS STAGE OF ALASKAN PORPHYRA (BANGIALES,RHODOPHYTA) SPECIES IN RESPONSE TO ENVIRONMENTAL VARIABLES1

Photosynthesis and respiration of three Alaskan Porphyra species, P. abbottiae V. Krishnam., P. pseudolinearis Ueda species complex (identified as P. “pseudolinearis” below), and P. torta V. Krishnam., were investigated under a range of environmental parameters. Photosynthesis versus irradiance (P–I) curves revealed that maximal photosynthesis (P_max), irradiance at maximal photosynthesis (I_max), and compensation irradiance (I_c) varied with salinity, temperature, and species. The P_max of Porphyra abbottiae conchocelis varied between 83 and 240 μmol O₂ · g dwt^?1 · h^?1 (where dwt indicates dry weight) at 30–140 μmol photons · m^?2 · s^?1 (I_max) depending on temperature. Higher irradiances resulted in photoinhibition. Maximal photosynthesis of the conchocelis of P. abbottiae occurred at 11°C, 60 μmol photons · m^?2·s^?1, and 30 psu (practical salinity units). The conchocelis of P. “pseudolinearis” and P. torta had similar P_max values but higher I_max values than those of P. abbottiae. The P_max of P. “pseudolinearis” conchocelis was 200–240 μmol O₂ · g dwt^?1 · h^?1 and for P. torta was 90–240 μmol O₂ · g dwt^?1 · h^?1. Maximal photosynthesis for P. “pseudolinearis” occurred at 7°C and 250 μmol photons · m^?2 · s^?1 at 30 psu, but P_max did not change much with temperature. Maximal photosynthesis for P. torta occurred at 15°C, 200 μmol photons · m^?2 · s^?1, and 30 psu. Photosynthesis rates for all species declined at salinities <25 or >35 psu. Estimated compensation irradiances (I_c) were relatively low (3–5 μmol · photons · m^?2 · s^?1) for intertidal macrophytes. Porphyra conchocelis had lower respiration rates at 7°C than at 11°C or 15°C. All three species exhibited minimal respiration rates at salinities between 25 and 35 psu.     

Studies of Elodea nuttallii grown under photorespiratory conditions. I. Photosynthetic characteristics

Abstract. The photosynthetic characteristics of Elodea nuttallii grown in wastewater in continuous flow reactors in a greenhouse were investigated. The diurnal changes in dissolved inorganic carbon (DIC), dissolved oxygen (DO) and pH were monitored. Photosynthesis removed both CO₂(aq) and HCO₃^? from the reactors. A stoichiometry of 1.19:1 was observed between HCO₃^? removal during photosynthesis and OH^? production during photosynthesis, consistent with theories regarding direct bicarbonate utilization. In laboratory experiments, the light compensation points (г_PPFD) were similar (31–35μmol m^?2 s^?1) to reported values for other macrophytes; however, the light saturation level was high (1100μmol m^?2 s^?1) and similar to values reported for aerial portions Of heterophyllous macrophytes. The kinetics of photosynthetic oxygen evolution (K_m (CO₂) = 96mmol m^?3; V_max= 133mmol g^?1 Chl h^?1) and the CO₂ compensation point (г= 44cm³ m^?3) suggested an adaptive, low photorespiratory state in response to low carbon concentrations. Photosynthetic V_max values were slightly, but significantly higher (P 0.001) at pH 8.0 compared to pH 4.5. While CO₂ utilization at pH 8 could account for most of the observed phototsynthetic rates, an HCO₃^? component was present, suggesting two separate transport systems for HCO₃^? and CO₂(aq) in E. nuttallii. The activity of RUBISCO (160.3 mmol g^?1 Chl h^?1 was one of the highest reported values for aquatic macrophytes. Compared to RUBISCO, we observed lower activities of the β-carboxylating enzymes phopho enolpyruvate carboyxlase (PEPcase), 24.1 mmol g^?1 Chl h^?1; phosphor enol pyruvate carboxykinase (PEPCKase), 14 mmol g^?1 Chl h^?1. This suggests that the potential light-independent fixation of carbon in E. nuttallii was much less than RUBISCO-dependent fixation. The RUBISCO/PEPcase ratio was 6.6, indicating that E. nuttallii was similar to Myriophyllum sp. in possessing a physiological adaptation to low CO₂ levels which is hypothesized to include carbonic anhydrase (CA) and an active transport system for HCO₃^?. CA levels were surprisingly low in E. nuttallii (14.2 EUmg Chl^?).     

Adaptations of tropical marine microphytobenthic assemblages along a gradient of light and nutrient availability in Suva Lagoon,Fiji

How are microphytobenthic biofilms adapted to the high incident irradiances and temperatures, low inorganic nutrient concentrations and high desiccation stresses on intertidal flats present in tropical environments? This study investigated biofilms subject to different environmental conditions in a range of tropical sites in Suva lagoon, Fiji. PAM fluorescence was used to measure photophysiological responses to the light climate. Biofilm colloidal carbohydrate, extracellular polymeric substances (EPS) and low molecular weight (MW) carbohydrate concentrations and diel carbohydrate production patterns were measured. Average biomass (Chl a) ranged from 15 to 36?mg?m^?2, and was highest in seagrass bed sediments, but biomass was not correlated with water column or sediment porewater nutrient concentrations. Biofilm photophysiology differed significantly along a combined gradient of light and nutrient availability, with F _v/F _m, relative ETR_max and E _k of biofilms highest in mangrove and intertidal main island sites and lowest in subtidal coral reef flats. Subtidal biofilms showed photoinhibition at irradiances > 1000?µmol?m^?2. Significant correlations between Chl a and colloidal carbohydrate concentrations were present (except on intertidal sandflats), and tropical biofilms had higher ratios of colloidal carbohydrate and EPS to Chl a than temperate estuarine biofilms, probably due to a combination of high irradiance and low nutrient availability leading to the production of excess photoassimilates. The percentage of EPS present in the colloidal fraction was highest in coral sand biofilms (42%), which had the lowest nutrient concentrations, compared with other sites (25–32%). Intertidal biofilms predominantly consisted of large motile taxa and showed strong rhythms of vertical migration. During tidal emersion, high sediment temperatures (41?°C), irradiance (>2300?µmol?m^?2?s^?1) and salinity (49‰) stimulated downward migration. In silty sediments, migration resulted in a reduction in photosynthetic activity during the midday period but, in sands with high light penetration (to a depth of > 1700?µm), high production rates of EPS (18.2?µg carbo. µg Chl a^?1 h^?1) and low MW carbohydrate exudates (40.2?µg carbo. µg Chl a^?1 h^?1) occurred. Vertical migration, high E _k and high rates of photoassimilate dumping are all adaptations to living in the tropical intertidal zone. Seagrass and reef flat biofilms consisted of a diverse non-migratory flora of motile and non-motile taxa that were not subject to such extreme temperature and irradiance conditions. Low values of photosynthetic parameters and high colloidal and EPS content indicated that these biofilms were nutrient-limited.     

Redistribution of phosphate deposits in the algaScenedesmus quadricauda deprived of exogenous phosphate—an ultra-cytochemical study

Summary The green algaScenedesmus quadricauda (Turp.) Bréb. was cultivated in the presence or absence of orthophosphate and synchronized daughter or mother cells were cytochemically stained. Forin situ capturing of water soluble phosphates Ca²⁺ and Mg²⁺ ions were added to the ice-cold glutaraldehyde fixative to form a polymeric metal-phosphate complex which was equivalent to the energy-rich condensed polyphosphates in staining by alkaline lead acetate. The X-ray microanalysis of the extensive stained deposits proved the presence of phosphorus. In orthophosphate-supplied daughter cells cytoplasmic vacuoles contained round stained bodies; a layer of phosphate-containing paracrystals encompassing some starch grains and a fine stained layer delineating the chloroplast envelope were also observed. In the equivalent mother cells only the material inside theloculi of stacked thylakoids was stained. In orthophosphate starved daughter cells filamentous phosphate-containing paracrystals filled extensive cytoplasmic vacuoles. A stained layer covered the chloroplast envelope and continuous stained layers appeared inside theloculi of stacked thylakoids. Mother cells that develop from these daughter cells were filled with starch grains and showed only peripheral stained deposits. The results are compared with the biochemical evidence of phosphate turnover in algal cells.Abbreviations ADP adenosine diphosphate - ATP adenosine triphosphate - ATPase adenosine triphosphatase - EDAX energy dispersive analysis of X-rays - Pi orthophosphate - PPi pyrophosphate - PP polyphosphate - PhAR photosynthetic active radiation - TCA trichloroacetic acid